The Refining the Shortest Paths (RSP) toolkit is a method for analyzing the fine-scale movements of aquatic animals tracked with passive acoustic telemetry in estuarine environments, that accounts for the surrounding land masses. The animal movements between detections are recreated to have occurred exclusively in water and the utilization distribution areas are limited by the land contours, providing realistic estimations of space use. The method can be divided into two main steps:
Depending on the research questions being addressed the utilization distribution areas can be calculated for the entire monitoring periods, or in fine-scale according to fixed temporal intervals (in hours). Tracked animals are assigned to specific biological groups (different species, different sexes from a same species, etc.) prior to analysis, and the RSP calculates the ammounts of inter-group overlap in space and time between all groups monitored. This approach allows spatial ecologists to use the outputs from such fine-scale space use models (areas of use, or between-group overlap frequencies) as input for further statistical analysis.
The RSP package can be downloaded using the following comand line:
install.packages("devtools")
library("devtools")
install_github("YuriNiella/RSP")
I think this first section might belong better in the README. Not sure if we had discussed this before!
Analysing acoustic telemetry data requires initial filtering to exclude misleading data (e.g. false detections, detections prior to release). To overcome this issue and ensure reliable results, the RSP toolkit operates in close relationship with the actel R package, which is filters and invalidates flawed detections. Before getting started with RSP, you will have to download actel and filter your acoustic data. Please click here for more information and to download actel.
To start using RSP, you can run the simplest actel analysis with the function explore. You can find more about how organize your data and run this preliminary analysis in actel’s manual pages (run ?actel after loading the library).
It is important that you save the output of the actel function, so you can later on use it to calculate the RSP. e.g.:
library(actel)
filtered_data <- explore(tz = "Europe/Sydney")
After filtering your acoustic data, you will need a raster file from your study area defining the water and land limits. This file will be used for estimating the shortest paths in water between consecutive acoustic detections, using a least-cost analysis of constrained random walks. The values of the raster cells must comprise zeros (water) or ones (land). Depending on the size of your study area, a resolution of 0.0001 (latitude ° x longitude °) is suggested for more accurate estimations (especially for sites with very narrow channels). Please see the following steps for generating and exporting a raster file from you study site in the QGIS software:
library(raster)
LakeMacquarie <- raster("Lake_Macquarie.grd") # Import the raster file exported from QGIS
plot(LakeMacquarie) # Plot raster from the study area
If the river channels look distorted go back to step 4 and choose a lower Cellsize. Please be aware* that increasing the raster resolution too much will require higher computational costs and may cause R to crash.
Now that your acoustic detections have been filtered and that you have a raster file with good resolution from your study area (exported into your working directory), you are all set to get started with RSP.
The runRSP() function is used to recreate the shortest paths between pairs of acoustic detections. The detection data, station coordinates and the group of each fish is passed on to RSP automatically by actel through the argument input. You must also include the name of the .grid file you created above in the argument base.raster. e.g.:
library(RSP)
rsp.results <- runRSP(input = filtered_data, base.raster = "Lake_Macquarie.grd")
A transition layer object is calculated using the raster file to estimate these paths exclusively in the water. The raster is automatically imported during the analysis through the argument base.raster = "name_of_your_file.grd". Because this step can take some time depending on the size of your study area and the size of the raster cells, the transition layer will be saved for future re-analysis in case the detection data changes. When running the analysis again the following message is shown:
M: Reusing transition layer calculated on 2019-11-30 13:48:19.
If you want to calculate a new transition layer, run rmTransition() before re-starting the analysis.
The detection ranges of each listening station are also taken into account in the runRSP(). These will be used as the location errors for the dBBMM when calculating UD areas. A ‘Range’ column can be included in the spatial.csv file for specifying the detection ranges (in meters) for each acoustic station if these are known. If the ‘Range’ column is not found, a default detection range of 500 m is automatically considered for each receiver with the warning:
Warning: Could not find a 'Range' column in the spatial data; assuming a range of 500 metres for each receiver.
Note: - The ‘Range’ column must already be present in the spatial.csv file when you run the explore() function for it to be incorporated in the analysis.
While animals move between a pair of consecutive acoustic detections there is some uncertainty regarding the trajectory taken, which increases proportionally to the time taken to from one place to another. Consecutive detections longer than 24 hours apart are thus broken by the runRSP() into separate ‘tracks’. This avoids the estimation of unrealistic behaviour when the animals do not get detected in any array for exceedingly long periods of time. Detections that occur totally isolated (e.g. more than 24-h before or after any other detection) are automatically excluded from analysis. The runRSP() will return the percentage of raw detections that can be used for refining the shortest paths when the analysis is finished:
M: Percentage of detections valid for RSP: 99.8%
Pairs of detections can occur either at the same receiver or at different receivers. For consecutive detections on different receivers, estimated positions are added according to a fixed distance argument in meters (250 m by default). On the other hand, if a fish is detected consecutively at the same station (with a time interval greater than the stipulated at the time.lapse argument), then estimated positions are added at that receiver location, over intervals of approximately [time.lapse] minutes. E.g. if a fish is detected at a station twice with a 22 minute interval, and time.lapse is set to 10, two estimated detections will be included.
While moving away from the first detection, the position errors gradually increase for each estimated position at a 5% rate of the distance. When the animal reaches half of the elapsed time/distance between the first and the second detection, the errors of estimated positions now gradually decrease as it approaches the second receiver where it got detected. This principle is used for both pairs of detections on different receivers, and for consecutive detections at the same station when the time difference is longer than time.lapse.
A: consecutive detections on the save receiver; B: consecutive detections on different receivers.
Please note how this animal was only detected on the same receivers during the first six tracks. Distances between consecutive RSP positions vary around the distance argument (250 m by default) as they depend on the estuary shape and the shortest distances between receivers. The same happens for the time lapses between consecutive RSP positions, which are around the time.lapse argument (10 min by default)
The dynamic Brownian Bridge Movement Model accounts for the speed at which animals move between consecutive detections to expand/contract the UD areas. Consequently, depending on your array configuration, estuary shape and species being tracked, you might find necessary to adjust the distance and time.lapse arguments to recreate the most plausible movement patterns of the monitored animals.
Here are some examples of the runRSP() output:
| Timestamp | Receiver | Transmitter | Error | Longitude | Latitude | Position | Track |
|---|---|---|---|---|---|---|---|
| 2018-03-07 00:43:49 | 125449 | R64K-4075 | 500 | 9.380188 | 56.5716 | Receiver | Track_3 |
| 2018-03-07 00:53:07 | NA | R64K-4075 | 512.5 | 9.380188 | 56.5716 | RSP | Track_3 |
| 2018-03-07 01:02:26 | NA | R64K-4075 | 512.5 | 9.380188 | 56.5716 | RSP | Track_3 |
| 2018-03-07 01:11:45 | 125449 | R64K-4075 | 550 | 9.380188 | 56.5716 | Receiver | Track_3 |
The Position column in this dataset identifies the two consecutive acoustic detections (Receiver) from this animal. We can notice that they occurred on the same Receiver (125449): the first on 2018-03-07 00:43:49 and the second on 2018-03-07 01:11:45 (approximately 30 minutes from each other). Because this time difference is longer than the default time.lapse (10 minutes), the runRSP() estimated the intermediate positions (RSP) by repeating the receiver Longitude and Latitude and changing the Error parameter (at a rate of 5% from the default distance argument of 250 meters = 12.5 meters).
| Timestamp | Receiver | Transmitter | Error | Longitude | Latitude | Position | Track |
|---|---|---|---|---|---|---|---|
| 2018-04-27 05:27:10 | 100474 | R64K-4125 | 500 | 9.921725 | 57.05595 | Receiver | Track_5 |
| 2018-04-27 05:35:17 | NA | R64K-4125 | 512.5 | 9.928500 | 57.05450 | RSP | Track_5 |
| 2018-04-27 05:43:24 | NA | R64K-4125 | 525 | 9.935500 | 57.05350 | RSP | Track_5 |
| 2018-04-27 05:51:32 | NA | R64K-4125 | 537.5 | 9.943500 | 57.05450 | RSP | Track_5 |
| 2018-04-27 05:59:39 | NA | R64K-4125 | 550 | 9.949500 | 57.05650 | RSP | Track_5 |
| 2018-04-27 06:07:47 | NA | R64K-4125 | 562.5 | 9.955500 | 57.05850 | RSP | Track_5 |
| 2018-04-27 06:15:54 | NA | R64K-4125 | 575 | 9.960500 | 57.06150 | RSP | Track_5 |
| 2018-04-27 06:24:01 | NA | R64K-4125 | 562.5 | 9.964500 | 57.06550 | RSP | Track_5 |
| 2018-04-27 06:32:09 | NA | R64K-4125 | 550 | 9.968500 | 57.06850 | RSP | Track_5 |
| 2018-04-27 06:40:16 | NA | R64K-4125 | 537.5 | 9.975500 | 57.07050 | RSP | Track_5 |
| 2018-04-27 06:48:24 | NA | R64K-4125 | 525 | 9.981500 | 57.07250 | RSP | Track_5 |
| 2018-04-27 06:56:31 | NA | R64K-4125 | 512.5 | 9.986500 | 57.07450 | RSP | Track_5 |
| 2018-04-27 07:04:39 | 107527 | R64K-4125 | 500 | 9.992500 | 57.07650 | Receiver | Track_5 |
Here the animal was detected first at the Receiver 100474 on 2018-04-27 05:27:10, and then at the Receiver 107527 on 2018-04-27 07:04:39. The runRSP() now calculated the shortest in-water path between receivers, and we can see how the Error of added locations increased up to half-way, (575 meters on 2018-04-27 06:15:54), and then decreased back to 500 as the track approached the second receiver.
We can use plotDistances() to compare the total distances travelled by each animal calculated using only the receiver locations and or also including the RSP estimations:
The plotDetections() shows the total number of receiver and estimated positions for each tracked animal:
You can also plot all tracks from a particular animal using plotRSP():
# Plot tracks only with receiver locations:
plotRSP(input = rsp.results, tag = "R64K-4075", display = "Receiver", type = "lines")
# Plot tracks with RSP estimations:
plotRSP(input = rsp.results, tag = "R64K-4075", display = "RSP", type = "lines")
You can also set display = "Both" to plot both track options on a single plot.
After estimating the in-water shortest paths, we can now use the output from runRSP() to calculate UD areas with the dynBBMM() function. Here you will need to know the UTM zone of your study site and specify it using the argument UTM.zone. By default, the analysis will run for all transmitters detected, but you can determine also which transmitters you would like to include using tags. As mentioned before, UD areas can be calculated with either of the following temporal resolutions:
This option calculates a series of dBBMM for each animal track from all the groups monitored. The breaks argument defines for which contours the areas of use should be calculated, which by default are the 50% and 95% (i.e. breaks = c(.5, .95)).
Track quality checks are performed to ensure that only good tracks which allow the dBBMM to converge are included in the analysis. This is an example of the returned messages from dynBBMM():
> group.dbbmm.results <- dynBBMM(input = rsp.results, UTM.zone = 56, breaks = c(0.5, 0.95))
M: Preparing data to apply dBBMM.
M: No specific transmitters selected. All the data will be used for analysis.
Warning: 8 track(s) in group Bream have less than eight detections and will not be used.
Warning: 5 track(s) in group Luderick have less than eight detections and will not be used.
Warning: 1 individual detections were removed in group Tarwhine due to simultaneous detections at two receivers.
Warning: 2 track(s) in group Tarwhine have less than eight detections and will not be used.
M: In total, 57 detections were excluded as they failed the track quality checks.
After calculating UDs, the land areas are excluded so that the final results represent only in-water areas of use. The overall overlap between each group monitored is also calculated.
M: Subtracting land areas from output.
M: Calculating overlaps between groups.
M: Storing final results.
The results of the dBBMM are saved in the $track.areas object, as a list of data frames for each group analysed:
| Track | Start | Stop | Area.5 | Area.95 | Time.lapse.min |
|---|---|---|---|---|---|
| R64K-4075_Track_3 | 2018-03-07 00:41:10 | 2018-03-07 08:20:02 | 20 | 49 | 458.87 |
| R64K-4075_Track_7 | 2018-04-24 11:40:56 | 2018-04-26 01:00:13 | 5817 | 17245 | 2239.28 |
| R64K-4075_Track_8 | 2018-08-20 11:56:47 | 2018-08-23 19:21:20 | 10609 | 36115 | 4764.55 |
| R64K-4125_Track_2 | 2018-04-21 13:13:24 | 2018-04-22 08:45:30 | 73 | 322 | 1172.10 |
| R64K-4125_Track_3 | 2018-04-25 11:44:05 | 2018-04-28 14:10:14 | 1469 | 16504 | 4466.15 |
| R64K-4125_Track_4 | 2018-08-23 12:16:57 | 2018-08-25 15:08:31 | 5875 | 15937 | 3051.57 |
| R64K-4125_Track_5 | 2018-08-27 08:44:04 | 2018-08-28 16:29:47 | 51 | 175 | 1905.72 |
| R64K-4125_Track_6 | 2018-08-30 17:26:41 | 2018-08-31 15:43:46 | 80 | 418 | 1337.08 |
| R64K-4128_Track_1 | 2018-04-13 09:44:22 | 2018-04-13 10:15:42 | 50 | 168 | 31.33 |
| R64K-4128_Track_3 | 2018-04-28 16:29:07 | 2018-04-29 15:25:16 | 2132 | 4793 | 1376.15 |
| R64K-4128_Track_4 | 2018-07-03 05:29:38 | 2018-07-05 13:29:06 | 3954 | 16292 | 3359.47 |
| R64K-4128_Track_5 | 2018-07-08 05:01:36 | 2018-07-08 22:43:43 | 478 | 2002 | 1062.12 |
| R64K-4138_Track_2 | 2018-03-07 23:39:11 | 2018-03-08 00:57:45 | 21 | 53 | 78.57 |
| R64K-4138_Track_6 | 2018-04-18 10:49:47 | 2018-04-20 12:38:33 | 4404 | 16396 | 2988.77 |
| R64K-4138_Track_7 | 2018-07-14 22:10:35 | 2018-07-18 18:33:43 | 7110 | 36902 | 5543.13 |
Each Track is named after the transmitter and the corresponding track name and both Start and Stop timestamps are stored. Areas of use (by default Area.5 and Area.95) are saved in squared meters, together with the respective elapsed times in minutes (Time.lapse.min). Note that in this example for the animal R64K-4075 only the Track_3, Track_7 and Track_8 were used for calculating space use areas, as the remainder ones failed the quality checks.
You can use plotContours() to visualize any of the dBBMM calculated by specifying the group and track you want to plot:
plotContours(input = group.dbbmm.results, group = "R64K-4138", track = 'R64K-4138_Track_7', main = "R64K-4138_Track_7")
The levels argument can be used to specify which contours to plot.
plotContours(input = group.dbbmm.results, group = "Bream", track = 'A69-9002-10473_Track_08', main = "A69-9002-10473_Track_08", stations = TRUE)
If stations = TRUE the locations of receiver stations are added to the plot.
(REVIEW STOPPED HERE!)
Now that we calculated the areas of space-use within our study area for each group monitored, we can investigate the ammount of overall overlap between the different species stored in the $overlap.areas object:
group.dbbmm.results$overlap.areas$`0.5`
group.dbbmm.results$overlap.areas$`0.5`$absolute
| Bream | Luderick | Tarwhine | |
|---|---|---|---|
| Bream | NA | NA | 10 |
| Luderick | NA | NA | NA |
| Tarwhine | 10 | NA | NA |
group.dbbmm.results$overlap.areas$`0.5`$percentage
| Bream | Luderick | Tarwhine | |
|---|---|---|---|
| Bream | NA | NA | 0.5263158 |
| Luderick | NA | NA | NA |
| Tarwhine | 0.5263158 | NA | NA |
group.dbbmm.results$overlap.areas$`0.95`
group.dbbmm.results$overlap.areas$`0.95`$absolute
| Bream | Luderick | Tarwhine | |
|---|---|---|---|
| Bream | NA | 2556 | 1962 |
| Luderick | 2556 | NA | 1205 |
| Tarwhine | 1962 | 1205 | NA |
group.dbbmm.results$overlap.areas$`0.95`$percentage
| Bream | Luderick | Tarwhine | |
|---|---|---|---|
| Bream | NA | 0.6336143 | 0.6254383 |
| Luderick | 0.6336143 | NA | 0.3841250 |
| Tarwhine | 0.6254383 | 0.3841250 | NA |
Please note the overlaps are calculated for the contours defined by breaks in dynBBMM(), and returned both in absolute values (squared meters) and percentage matrices. For example, we can see in the last table that Bream and Luderick were the groups with higher overall overlap of 63.36% at the 95% level, whereas Tarwhine and Luderick only had a 38.41% overlap at the 95% contour. To see exactly where space use overlaps occurred you can use plotOverlap():
plotOverlap(input = group.dbbmm.results, stations = FALSE, level = .95, store = TRUE)
dBBMMs can also be calculated according to a moving temporal window. This allows investigating how the space-use overlap between the different groups varied during the study period. It is useful for assessing the influence of environmental parameters upon space-use of different groups tracked within the study area. The same dynBBMM() function is used, but here the argument timeframe has to be defined in hours as the temporal window. The total tracking period will be divided into timeslots, and dBBMMs calculated for each group monitored (for each timeslot). Overlapping areas are now calculated for each timeslot and the corresponding metadata stored in the $timeslots object:
time.dbbmm.results <- dynBBMM(input = rsp.results, UTM.zone = 56, breaks = c(0.5, 0.95), timeframe = 12) # 12-h timeslots
head(time.dbbmm.results$timeslots)
| slot | start | stop | Bream | Luderick | Tarwhine |
|---|---|---|---|---|---|
| 1 | 2013-09-02 00:00:00 | 2013-09-02 12:00:00 | FALSE | FALSE | FALSE |
| 2 | 2013-09-02 12:00:00 | 2013-09-03 00:00:00 | FALSE | TRUE | FALSE |
| 3 | 2013-09-03 00:00:00 | 2013-09-03 12:00:00 | FALSE | TRUE | TRUE |
| 4 | 2013-09-03 12:00:00 | 2013-09-04 00:00:00 | FALSE | FALSE | TRUE |
| 5 | 2013-09-04 00:00:00 | 2013-09-04 12:00:00 | TRUE | FALSE | TRUE |
| 6 | 2013-09-04 12:00:00 | 2013-09-05 00:00:00 | TRUE | FALSE | TRUE |
In the example above we can notice that in the first timeslot none of the groups were detected, whereas Luderick group got detected in slot 2 and both Luderick and Tarwhine were detected on the first 12-h of 2013-09-03 (slot 3). The $track.areas object for each tracked group will now have a first column named Slot, which identifies the timeslot for each of the dBBMM calculated:
head(time.dbbmm.results$track.areas)?
what was the command to get the table below exactly?
| Slot | Track | Start | Stop | Area.5 | Area.95 | Time.lapse.min |
|---|---|---|---|---|---|---|
| 486 | A69-9002-10474_Track_1 | 2014-05-02 12:11:54 | 2014-05-02 23:54:41 | 249 | 962 | 702.7833 |
| 486 | A69-9002-10480_Track_1 | 2014-05-02 12:21:36 | 2014-05-02 23:37:18 | 279 | 1215 | 675.7000 |
| 487 | A69-9002-10474_Track_1 | 2014-05-03 00:12:19 | 2014-05-03 11:42:37 | 238 | 897 | 690.3000 |
| 487 | A69-9002-10480_Track_1 | 2014-05-03 01:18:26 | 2014-05-03 11:56:30 | 1170 | 4516 | 638.0667 |
| 488 | A69-9002-10474_Track_1 | 2014-05-03 12:11:19 | 2014-05-03 23:56:57 | 274 | 1051 | 705.6333 |
| 488 | A69-9002-10480_Track_1 | 2014-05-03 12:26:10 | 2014-05-03 23:48:24 | 345 | 1396 | 682.2333 |
Here we can see that two animals were detected for this particular group between Slot 486 and Slot 488: the transmitters A69-9002-10474 and A69-9002-10480. The Time.lapse.min will now be very similar as its maximum elapsed time is limited by the timeframe argument, and it depends on the times of first and last locations for each animal.
The following command line can help you assess if any other group got detected during this same timeslot:
> time.dbbmm.results$timeslots[486, ]
slot start stop Bream Luderick Tarwhine
486 486 2014-05-02 12:00:00 2014-05-03 00:00:00 TRUE TRUE FALSE
Yes, both Bream and Luderick were detected between 2014-05-02 12:00:00 and 2014-05-03 00:00:00. We can inspect whether the two groups overlapped or not:
> time.dbbmm.results$overlap.areas$`0.95`$percentage$`486`
Bream Luderick Tarwhine
Bream NA 0.8247734 NA
Luderick 0.8247734 NA NA
Tarwhine NA NA NA
This shows that the two groups had an overlap of 82.47% at the 95% dBBMM contour during this particular timeslot. We can now see exactly where the overlap occurred by plotting the space-use models using:
# plotContours() for plotting the individual dBBMM:
plotContours(input = time.dbbmm.results, track = "A69-9002-10474_Track_1", group = "Bream", timeslot = 486, stations = TRUE, main = "A69-9002-10474 (Bream - slot 486)")
plotContours(input = time.dbbmm.results, track = "A69-9002-10480_Track_1", group = "Bream", timeslot = 486, stations = TRUE, main = "A69-9002-10480 (Bream - slot 486)")
plotContours(input = time.dbbmm.results, track = "A69-9002-10481_Track_1", group = "Luderick", timeslot = 486, stations = TRUE, main = "A69-9002-10481 (Luderick - slot 486)")
# plotOverlap() to investigate the overlap areas:
plotOverlap(input = time.dbbmm.results, store = TRUE, stations = FALSE, timeslot = 486)
Approximately 1-month of fine-scale (12-h) space use of Bream and Luderick at the 95% level